Phylogenetic-Signal Dissection of Nuclear Housekeeping Genes Supports the Paraphyly of Sponges and the Monophyly of Eumetazoa

The relationships at the base of the metazoan tree have been difficult to robustly resolve, and there are several different hypotheses regarding the interrelationships among sponges, cnidarians, ctenophores, placozoans, and bilaterians, with each hypothesis having different implications for the body plan of the last common ancestor of animals and the paleoecology of the late Precambrian. We have sequenced seven nuclear housekeeping genes from 17 new sponges, bringing the total to 29 species analyzed, including multiple representatives of the Demospongiae, Calcarea, Hexactinellida, and Homoscleromorpha, and analyzed a data set also including six nonmetazoan outgroups and 36 eumetazoans using a variety of phylogenetic methods and evolutionary models. We used leaf stability to identify rogue taxa and investigate their effect on the support of the nodes in our trees, and we identified clades most likely to represent phylogenetic artifacts through the comparison of trees derived using different methods (and models) and through site-stripping analyses. Further, we investigated compositional heterogeneity and tested whether amino acid composition bias affected our results. Finally, we used Bayes factors to compare our results against previously published phylogenies. All our maximum likelihood (ML) and Bayesian analyses find sponges to be paraphyletic, with all analyses finding three extant paraphyletic sponge lineages, Demospongiae plus Hexactinellida, Calcarea, and Homoscleromorpha. All but one of our ML and Bayesian analyses support the monophyly of Eumetazoa (here Cnidaria þ Bilateria) and a sister group relationship between Placozoa (here Trichoplax adhaerens) and Eumetazoa. Bayes factors invariably provide decisive support in favor of poriferan paraphyly when compared against either a sister group relationship between Porifera and Cnidaria or with a monophyletic Porifera with respect to a monophyletic Eumetazoa. Although we were able to recover sponge monophyly using our data set, this was only possible under unrealistic evolutionary models, if poorly performing phylogenetic methods were used, or in situations where the potential for the generation of tree reconstruction artifacts was artificially exacerbated. Everything considered, our data set does not provide any support for a monophyletic Diploblastica (here Placozoa þ Cnidaria þ Porifera) and suggests that a monophyletic Porifera may be better seen as a phylogenetic artifact

Phylogenetic-Signal Dissection of Nuclear Housekeeping Genes Supports the Paraphyly of Sponges and the Monophyly of Eumetazoa

The relationships at the base of the metazoan tree have been difficult to robustly resolve, and there are several different hypotheses regarding the interrelationships among sponges, cnidarians, ctenophores, placozoans, and bilaterians, with each hypothesis having different implications for the body plan of the last common ancestor of animals and the paleoecology of the late Precambrian. We have sequenced seven nuclear housekeeping genes from 17 new sponges, bringing the total to 29 species analyzed, including multiple representatives of the Demospongiae, Calcarea, Hexactinellida, and Homoscleromorpha, and analyzed a data set also including six nonmetazoan outgroups and 36 eumetazoans using a variety of phylogenetic methods and evolutionary models. We used leaf stability to identify rogue taxa and investigate their effect on the support of the nodes in our trees, and we identified clades most likely to represent phylogenetic artifacts through the comparison of trees derived using different methods (and models) and through site-stripping analyses. Further, we investigated compositional heterogeneity and tested whether amino acid composition bias affected our results. Finally, we used Bayes factors to compare our results against previously published phylogenies. All our maximum likelihood (ML) and Bayesian analyses find sponges to be paraphyletic, with all analyses finding three extant paraphyletic sponge lineages, Demospongiae plus Hexactinellida, Calcarea, and Homoscleromorpha. All but one of our ML and Bayesian analyses support the monophyly of Eumetazoa (here Cnidaria þ Bilateria) and a sister group relationship between Placozoa (here Trichoplax adhaerens) and Eumetazoa. Bayes factors invariably provide decisive support in favor of poriferan paraphyly when compared against either a sister group relationship between Porifera and Cnidaria or with a monophyletic Porifera with respect to a monophyletic Eumetazoa. Although we were able to recover sponge monophyly using our data set, this was only possible under unrealistic evolutionary models, if poorly performing phylogenetic methods were used, or in situations where the potential for the generation of tree reconstruction artifacts was artificially exacerbated. Everything considered, our data set does not provide any support for a monophyletic Diploblastica (here Placozoa þ Cnidaria þ Porifera) and suggests that a monophyletic Porifera may be better seen as a phylogenetic artifact

Tapped Out: Threats to the Human Right to Water in the Urban United States

In the United States today, the goal of universal water service is slipping out of reach. Water costs are rising across the country, forcing many individuals to forgo running water or sanitation, or to sacrifice other essential human rights. The fixed costs of water systems have increased in recent years, driven in part by underinvestment in infrastructure. In many cities, this has been exacerbated by population shifts and the economic downturn. In this era of increasing costs and limited financial resources, water providers struggle to balance the competing priorities of modernization and universal access. This report, researched and written by students of Georgetown Law’s Human Rights Institute in the winter of 2013, details the causes, effects, and solutions to the affordability crisis affecting water in the urban United States

Oxygen and animals in Earth history

Earth and Planetary Science

A Basin Redox Transect at the Dawn of Animal Life

Multiple eukaryotic clades make their first appearance in the fossil record between ~810 and 715 Ma. Molecular clock studies suggest that the origin of animal multicellularity may have been part of this broader eukaryotic radiation. Animals require oxygen to fuel their metabolism, and low oxygen levels have been hypothesized to account for the temporal lag between metazoan origins and the Cambrian radiation of large, ecologically diverse animals. Here, paleoredox conditions were investigated in the Fifteenmile Group, Ogilvie Mountains, Yukon, Canada, which hosts an 811 Ma ash horizon and spans the temporal window that captures the inferred origin and early evolution of animals. Iron-based redox proxies, redox-sensitive trace elements, organic carbon percentages and pyrite sulfur isotopes were analyzed in seven stratigraphic sections along two parallel basin transects. These data suggest that for this basin, oxygenated shelf waters overlay generally anoxic deeper waters. The anoxic water column was dominantly ferruginous, but brief periods of euxinia likely occurred. These oscillations coincide with changes in total organic carbon, suggesting euxinia was primarily driven by increased organic carbon loading. Overall, these data are consistent with proposed quantitative constraints on Proterozoic atmospheric oxygen being greater than 1% of modern levels, but less than present levels. Comparing these oxygen levels against the likely oxygen requirements of the earliest animals, both theoretical considerations and the ecology of modern oxygen-deficient settings suggest that the inferred oxygen levels in the mixed layer would not have been prohibitive to the presence of sponges, eumetazoans or bilaterians. Thus the evolution of the earliest animals was probably not limited by the low absolute oxygen levels that may have characterized Neoproterozoic oceans, although these inferred levels would constrain animals to very small sizes and low metabolic rates.Earth and Planetary SciencesOrganismic and Evolutionary Biolog

The mitochondrial genome of the hexactinellid sponge Aphrocallistes vastus: Evidence for programmed translational frameshifting

<p>Abstract</p> <p>Background</p> <p>Mitochondrial genomes (mtDNA) of numerous sponges have been sequenced as part of an ongoing effort to resolve the class-level phylogeny of the Porifera, as well as to place the various lower metazoan groups on the animal-kingdom tree. Most recently, the partial mtDNA of two glass sponges, class Hexactinellida, were reported. While previous phylogenetic estimations based on these data remain uncertain due to insufficient taxon sampling and accelerated rates of evolution, the mtDNA molecules themselves reveal interesting traits that may be unique to hexactinellids. Here we determined the first complete mitochondrial genome of a hexactinellid sponge, <it>Aphrocallistes vastus</it>, and compared it to published poriferan mtDNAs to further describe characteristics specific to hexactinellid and other sponge mitochondrial genomes.</p> <p>Results</p> <p>The <it>A. vastus </it>mtDNA consisted of a 17,427 base pair circular molecule containing thirteen protein-coding genes, divergent large and small subunit ribosomal RNAs, and a reduced set of 18 tRNAs. The <it>A. vastus </it>mtDNA showed a typical hexactinellid nucleotide composition and shared a large synteny with the other sequenced glass sponge mtDNAs. It also contained an unidentified open reading frame and large intergenic space region. Two frameshifts, in the <it>cox3 </it>and <it>nad6 </it>genes, were not corrected by RNA editing, but rather possessed identical shift sites marked by the extremely rare tryptophan codon (UGG) followed by the common glycine codon (GGA) in the +1 frame.</p> <p>Conclusion</p> <p>Hexactinellid mtDNAs have shown similar trends in gene content, nucleotide composition, and codon usage, and have retained a large gene syntenty. Analysis of the mtDNA of <it>A. vastus </it>has provided evidence diagnostic for +1 programmed translational frameshifting, a phenomenon disparately reported throughout the animal kingdom, but present in the hexactinellid mtDNAs that have been sequenced to date.</p

The Ecological Physiology of Earth's Second Oxygen Revolution

Living animals display a variety of morphological, physiological, and biochemical characters that enable them to live in low-oxygen environments. These features and the organisms that have evolved them are distributed in a regular pattern across dioxygen (O2) gradients associated with modern oxygen minimum zones. This distribution provides a template for interpreting the stratigraphic covariance between inferred Ediacaran-Cambrian oxygenation and early animal diversification. Although Cambrian oxygen must have reached 10--20% of modern levels, sufficient to support the animal diversity recorded by fossils, it may not have been much higher than this. Today’s levels may have been approached only later in the Paleozoic Era. Nonetheless, Ediacaran-Cambrian oxygenation may have pushed surface environments across the low, but critical, physiological thresholds required for large, active animals, especially carnivores. Continued focus on the quantification of the partial pressure of oxygen (pO2) in the Proterozoic will provide the definitive tests of oxygen-based coevolutionary hypotheses.Earth and Planetary SciencesOrganismic and Evolutionary Biolog

Exploring the Role of Cities in Electrifying Passenger Transportation

Key Takeaways1. The electrification of passenger vehicles should be one part of a city’s transportation plan. Shifting from internal combustion engine vehicles to plug-in electric vehicles (PEVs) can improve urban air quality, reduce greenhouse gas emissions, and reduce energy consumption.2. Recent studies show that electric vehicle awareness is low even in mature markets; cities should promote electric vehicles to residents by leveraging existing promotional campaigns.3. Various financial and non-financial incentives can effectively encourage electric vehicle uptake, including: free, discounted, or preferential-location parking; free or reduced road and bridge tolls; and allowing electric vehicles to drive in bus or carpool lanes.4. Several cities are restricting or planning to restrict the access that internal combustion engine vehicles (ICEVs) have to certain areas. If these restrictions apply to most (or all) passenger ICEVs, they can promote PEV purchase and use in cities.5. Infrastructure development in cities should follow the same fundamental approach as that used outside of cities. The priority should be ensuring that PEV owners and prospective PEV buyers have access to charging at or near home. Workplace and public charging should be developed for those who cannot access charging at or near home.6. Cities should be strategic in their approach, first identifying the goals they want to achieve, and then exploring what steps they can take to meet these goals. The steps available will likely differ between cities due to the different ways in which roads, parking, and any other vehicle infrastructure is governed

Persistent global marine euxinia in the early Silurian

The second pulse of the Late Ordovician mass extinction occurred around the Hirnantian-Rhuddanian boundary (~444 Ma) and has been correlated with expanded marine anoxia lasting into the earliest Silurian. Characterization of the Hirnantian ocean anoxic event has focused on the onset of anoxia, with global reconstructions based on carbonate δ238U modeling. However, there have been limited attempts to quantify uncertainty in metal isotope mass balance approaches. Here, we probabilistically evaluate coupled metal isotopes and sedimentary archives to increase constraint. We present iron speciation, metal concentration, δ98Mo and δ238U measurements of Rhuddanian black shales from the Murzuq Basin, Libya. We evaluate these data (and published carbonate δ238U data) with a coupled stochastic mass balance model. Combined statistical analysis of metal isotopes and sedimentary sinks provides uncertainty-bounded constraints on the intensity of Hirnantian-Rhuddanian euxinia. This work extends the duration of anoxia to &gt;3 Myrs – notably longer than well-studied Mesozoic ocean anoxic events